Description of two new species of fig wasps (Chalcidoidea: Pteromalidae: Sycoryctinae) associated with Ficus benguetensis
←
→
Page content transcription
If your browser does not render page correctly, please read the page content below
Taiwania 63(2): 155-162, 2018
DOI: 10.6165/tai.2018.63.155
Description of two new species of fig wasps (Chalcidoidea: Pteromalidae:
Sycoryctinae) associated with Ficus benguetensis
Da-Mien WONG1, Anthony BAIN2, Lien-Siang CHOU1 and Shiuh-Feng SHIAO3,*
1. Institute of Ecology and Evolutionary Biology, College of Life Science, National Taiwan University, 1, Sec. 4, Roosevelt Rd.,
Taipei, 10617, Taiwan.
2. Department of Biological Sciences, National Sun Yat-sen University, 70 Lienhai Rd., Kaohsiung 80424, Taiwan.
3. Department of Entomology, National Taiwan University, 1, Sec. 4, Roosevelt Rd., Taipei, 10617, Taiwan.
*
Corresponding author’s tel. +886-02-33665530, fax. +886-02-27359990, email: sfshiao@ntu.edu.tw
(Manuscript received 4 April 2018; accepted 18 May 2018; online published 28 May 2018)
ABSTRACT: This study presents two new fig wasp species, Philotrypesis taida Wong and Shiao sp. nov. and Sycorycteridea
taipeiensis Wong and Shiao sp. nov., which were reared from Ficus benguetensis Merrill in Taiwan. They are the seventh and first
species belonging to the genera Philotrypesis and Sycorycteridea, respectively, to be reported in Taiwan. The holotypes and
paratypes were deposited in the Department of Entomology at National Taiwan University, Taiwan.
KEY WORDS: Chalcidoidea, Ficus, Fig wasps, Philotrypesis, Sycorycteridea, Taiwan.
INTRODUCTION galls. Males usually fight severely using their
mandibles for females soon after their emergence
Fig wasps belonging to the subfamily Sycoryctinae (Murray, 1987; Bean and Cook, 2001). Male
(Chalcidoidea: Pteromalidae) are highly diversified and Sycoryctinae exhibit diverse adaptations in their
widespread parasitoid species (Segar et al., 2012). morphology, including winged dispersers and wingless
Sycoryctine fig wasps are associated with all six fighters (Jousselin et al., 2004). Sycoryctines serve as a
subgenera and at least 15 sections in Ficus (Jiang et al., model organism in behavioral ecology research because
2006). Consequently, there is an estimated species of their fascinating behaviors and diverse morphologies
richness of 826 that is yet to be discovered in the Old (Moore et al., 2009). Despite the ecological importance
World (Segar et al., 2012). From a taxonomic of sycoryctine fig wasps as nonpollinating species
perspective, Sycoryctinae are currently placed in the associated with fig trees, they are poorly represented in
family Pteromalidae within the superfamily collections in Taiwan because of the special
Chalcidoidea (Rasplus et al., 1998). Sycoryctinae are requirements for collecting, preserving, and identifying
then divided into four tribes: (1) Apocryptini, them. Consequently, only eight species of sycoryctine
comprising two genera: Apocrypta Coquerel, 1855, and fig wasps have been identified in Taiwan (Chou and
Bouceka Kocak and Kemal, 2008; (2) Critogastrini, Wong, 1997; Chen et al., 1999).
comprising only one genus: Critogaster Mayr, 1885; (3) Two new fig wasp species belonging to the genera
Philotrypesini, comprising four genera: Dobunabaa Philotrypesis and Sycorycteridea, which are associated
Boucek, 1988, Philoverdance Priyadarsanan, 2000, with the fig tree Ficus benguetensis Merrill, are
Philotrypesis Forster, 1878, and Watshamiella Wiebes, reported in this study. They are the seventh and first
1981; and (4) Sycoryctini, comprising seven genera: species belonging to the genera Philotrypesis and
Adiyodiella Priyadarsanan, 2000, Arachonia Joseph, Sycorycteridea, respectively, reported in Taiwan.
1957, Parasycobia Abdurahiman and Joseph, 1967,
Sycorycteridea Abdurahiman and Joseph, 1967, MATERIALS AND METHODS
Sycoryctes Mayr, 1885, Sycoscapter Saunders, 1883,
and Sycoscapteridea Ashmead, 1904 (McLeish et al., Fig wasp samples associated with the fig tree F.
2012; Segar et al., 2012). benguetensis were collected on the National Taiwan
Female sycoryctines oviposit through the fig wall University (NTU) campus, Taipei from 2011 to 2013.
with the aid of their long ovipositors, and their larvae Additional populations were collected from Fuyang Eco
eventually kill their host wasps or spend their larval Park, Taipei and Hengchun Tropical Garden, Pingtung
stage inside the figs as plant-feeders (Compton et al., during 2014 to 2017 (Table 1). The sample figs were
2015). The feeding habits of sycoryctine larvae vary stored at room temperature in transparent plastic pots
considerably as they can be parasitoids, inquilines, or closed with mesh covers for 24 hours with their ostioles
obligate seed predators, among other types (Tzeng et al., facing upward. The wasps’ exit was confirmed by the
2008; Wang et al., 2014). Sycoryctine larvae mature in presence of a hole, which was chewed on the surface of
155
Taiwania Vol. 63, No. 2
Table 1: Collection point of Philotrypesis taida sp. nov. and Sycorycteridea taipeiensis sp. nov.
Date Coordinate Location Host (Ficus) Number of figs
2011-Oct-29 25°00'54.7"N; 121°32'11.8"E NTU Campus F. benguetensis 10
2012-Jul-26 25°00'54.7"N, 121°32'11.8"E NTU Campus F. benguetensis 15
2013-Aug-17 25°00'57.6"N, 121°33'27.0"E Fuyang Eco Park F. benguetensis 8
2014-Jul-20 25°00'57.6"N, 121°33'27.0"E Fuyang Eco Park F. benguetensis 10
2015-Jul-22 25°00'57.6"N, 121°33'27.0"E Fuyang Eco Park F. benguetensis 10
2017-May-20 21°57'45.0"N, 120°48'38.2"E Hengchun Tropical Garden F. benguetensis 15
2017-Jul-14 21°57'45.0"N, 120°48'38.2"E Hengchun Tropical Garden F. benguetensis 10
the figs. All the wasps that exited their natal figs were inhabiting Taiwan is provided (Table 2).
collected and stored in 75% ethanol. The figs were then Types: Holotype, ♀, TAIWAN: Taipei City: Daan
dissected and the wasps therein were also collected. District: NTU campus, 25°00'54.7"N; 121°32'11.8"E,
The collected wasps were dehydrated through an Alt. 0–10 m, October 29, 2011, D.M. Wong; Paratypes,
ethanol series (75%, 90%, and 100%) and critical-point 5♀ and 5♂, the same locality and data as holotype. All
dried (DO-11-R-269 CPD, Quorum Technologies Inc., specimens deposited in the Department of Entomology
UK) before being mounted on cards following Noyes at NTU.
(1982). The specimens were point-mounted on black Description: Female - Body length 2.0–2.2 mm.
cards and photographed to avoid the low contrast Body color variable. Head and antenna orange.
caused by a white background. Mandible bidentate. Mesosoma and metasoma usually
Morphological terminology followed Gibson (1997) brownish orange, sometimes metallic black. Coxae
and the Hymenoptera Anatomy Ontology (HAO) Portal concolorous with mesosoma. Wings hyaline. The 7th
(Yoder et al., 2010). Images were taken using a and 8th segment in ratio 3:1. Ovipositor sheath length
stereomicroscope (Leica Z16 APO, Wetzlar, Germany) 3.0–3.6 mm.
with a digital camera (Leica DFC490, Wetzlar, Head. Head width 0.4–0.5 mm (Fig. 1A). Antenna
Germany) and were processed using the Helicon Focus with 13 antennomeres, antennal formula: 11353 (Fig.
6.7.1. software (Helicon Soft Inc., Dominica) to 1B). Scape clavate 4× longer than goblet pedicel.
combine them into a single image with increased focal Funicular segments slightly longer than wide. Terminal
depth. Physical characteristics were measured using antennomere conspicuous. Face sculpture smooth.
ImageJ 1.45a software (National Institutes of Health, Antenna inserted at the bottom line of compound eyes.
USA). Specimen measurements were taken with Toruli apart, distance between toruli larger than
accuracy of 0.001 mm and were rounded to the nearest diameter of one torulus. Eye red, longer than gena.
0.1 mm. Holotype and paratype specimens were Clypeal margin flattened. Mandible bidentate.
deposited in the Department of Entomology at NTU. Mesosoma. Body length 2.0–2.2 mm (Fig. 1C);
Coxae concolorous with mesosoma. Hind tibia at its
TAXONOMIC TREATMENT distal end provided with apical spur and about 8× as long
as its own thickness (Fig. 1D). Wings length 1.3–1.5 mm
Key to species of Philotrypesis females commonly and finely pubescent (Fig. 1E). Postmarginal vein 3×
found in Taiwan longer than stigmal vein. Pronotum sculpture smooth
(Fig. 1F). Pronotum longer than high in lateral view.
1a. Body black; 2 anelli; mandibles 2 teeth ………………………….. Notauli complete. Mesoscutellum trapezoidal. Axilla
…………..………………….….… Philotrypesis taiwanensis Chen
1b. Body orange; 3 anelli; mandibles 2 or 3 teeth …………..……… 2 triangular and small. Scutellum round. Propendeum
2a. Mandible 2 teeth ……………………….……………………..… 3 transverse.
2b. Mandible 3 teeth ………………..… Philotrypesis emeryi Grandi Metasoma. Without petiole. The 7th and 8th segment
3a. Gaster VII as long as VIII (1:1) .... Philotrypesis okinavensis Ishii in ratio 3:1. Ovipositor sheath length 1.5× longer than
3b. Gaster VII longer than VIII (3:1) …... Philotrypesis taida sp. nov.
body.
Philotrypesis taida Wong & Shiao, sp. nov. Male - Head width 0.3–0.7 mm and covered long
臺大延腹小蜂 Figs. 1–2 seta (Fig. 2A). Head as long as wide across eye.
urn:lsid:zoobank.org:act:8EC5D142-54DC-4035-B41D-1F2FA9A54B20
Mandible length 0.1–0.4 mm. Eye black, one-fourth of
Diagnosis: The female of this genus can be head length. Antenna toruli situate in a shallow
recognized by its lengthened seventh and eighth depression above epistomal margin, with 1 keel in the
urotergites and its subquadrate narrow pronotum. The middle. Body orange, apterous and body length 2.8–3.6
antenna of this genus usually composed of 12–13 mm (Fig. 2B–C).
antennomeres (1–3 anelli), including an oval terminal Etymology: The specific name refers to the pinyin
flagellomere. Comparison of morphological (Chinese pronunciation) abbreviation of “National
characteristics in females between Philotrypesis spp. Taiwan University.”
156
June 2018 Wong et al.: New fig wasps associated with Ficus benguetensis
Fig. 1. Female Philotrypesis taida sp. nov. A: Head frontal view; B: Antenna; C: Lateral view; D: Hind leg; E: Wings; F: Dorsal view of
mesosoma. Scale bar = 0.2 mm.
157
Taiwania Vol. 63, No. 2
Fig. 2. Male Philotrypesis taida sp. nov. A: Head frontal view; B: Dorsal view of mesosoma; C: Lateral view. Scale bar = 0.2 mm.
Host: Reared from syconia of F. benguetensis. Table 2: Characteristics of four Philotrypesis species
inhabiting Taiwan
Biology: This species could be the parasitoids or
inquilines of other fig wasps associated with fig sections, Species P. P. P. P.
such as Conosycea, Ficus, Galoglychia, Sycidium, (Philotrypesis) taida taiwanensis okinavensis emeryi
Body color Orange Black Orange Orange
Sycocarpus, and Urostigma. Across eye Below eye Below eye Below eye
Distribution: Taiwan. According to host Antenna toruli
area area area area
specialization of fig wasps and the geographical Anelli 3 2 3 3
distribution of F. benguetensis, we assume that this Mandible teeth 2 2 2 3
Gaster VII : VIII 3:1 3:2 1:1 3:1
species is also distributed in Malesia and Ryukyu F. F. F. F.
islands. Host (Ficus)
benguetensis microcarpa microcarpa microcarpa
158
June 2018 Wong et al.: New fig wasps associated with Ficus benguetensis
Sycorycteridea taipeiensis Wong & Shiao, sp. nov. Distribution: Taiwan. According to host
臺北細尾小蜂 Figs. 3 & 4 specialization of fig wasps and the geographical
urn:lsid:zoobank.org:act:9C6EA98B-0E54-40DA-B995-87F8A8BDC5D4 distribution of F. benguetensis, we assume that this
Diagnosis: The female of this genus can be species is also distributed in Malesia and Ryukyu islands.
recognized by the long slender ovipositor and its sheaths,
slightly swollen at the apex, the whole length of the Table 3: Characteristics comparison between female
Sycorycteridea taipeiensis and S. keralensis
ovipositor being covered by the extended last tergite.
Tarsi distinctly shorter than tibia. Comparison of S. taipeiensis S. keralensis
morphological characteristics in females between Body color Yellow Metallic green
Sycorycteridea taipeiensis sp. nov. and Sycorycteridea Anellus 1 1
keralensis is provided (Table 3). Epistomal margin Protruded Protruded
Ovipositor : body 3:1 5:1
Types: Holotype ♀, TAIWAN: Taipei City: Daan Stigmal knob Not elongate Not elongate
District: NTU campus, 25°00'54.7"N; 121°32'11.8"E, Alt. Host (Ficus) F. benguetensis F. callosa
0–10 m, October 29, 2011, D.M. Wong; Paratypes, 5♀,
5♂, same locality and data as holotype. All specimens DISCUSSION
deposited in the Department of Entomology at NTU.
Description: Female - Body length 1.0–1.5 mm. Fig wasps are pantropically distributed and play a
Body color variable. Antenna yellow. Head and pertinent role within tropical plant communities (Bain
mesosoma yellow to metallic black. Metasoma usually et al., 2015). Fig wasps and their host fig trees have
yellow, sometimes metallic black. Coxae concolorous long been used as an example in studies on interspecific
with mesosoma. Wings with short and sparse pilosity. interactions. These fascinating interactions include
Ovipositor sheath length 3.0–3.2 mm. classic obligate mutualism to antagonistic parasitism.
Head. Head width 0.2–0.3 mm (Fig. 3A). Antenna These trophic relationships of the figs and wasps
with 12 antennomeres, antennal formula: 11163 (Fig. sometimes include up to 20 taxa on a single fig species
3B). Scape clavate 3× longer than goblet pedicel. (Chen et al., 1999). Because plant–herbivore–parasitoid
Funicular segments slightly longer than wide. Terminal associations are a major component of terrestrial
antennomere conspicuous. Antenna inserted at the biodiversity, intensive taxonomical research is hence
bottom line of compound eyes. Toruli contiguous, pertinent for understanding how fig wasp communities
distance between toruli smaller than diameter of one interact and evolve.
torulus. Ecdysial cleavage conspicuous. Eye red, longer Although some examples of host switching have
than gena. Face sculpture smooth. Clypeal margin been found, fig wasps belonging to Sycoryctinae are the
protruded. Mandible bidentate. largest nonpollinating wasps conservatively associated
Mesosoma. Body length 1.0–1.5 mm (Fig. 3C); with fig trees in Old World regions (Segar et al., 2012).
Coxae concolorous with mesosoma. Hind tibia at its Despite being geographically widespread, the taxonomy
distal end provided with apical spur and about 6× as long of sycoryctine fig wasps is poorly understood and is less
as its own thickness (Fig. 3D). Tarsi distinctly shorter advanced than that of pollinating fig wasp species.
than tibia. Wings length 0.9–1.0 mm with short and Sycoryctine fig wasps inhabiting the Indomalayan region
sparse pilosity (Fig. 3E). Postmarginal vein 2.5× longer have received less taxonomical attention compared with
than stigmal vein. Pronotum sculpture smooth (Fig. 3F). those from other biogeographical realms, such as the
Pronotum longer than high in lateral view. Notauli Afrotropics and Neotropics and Australasia (Bouček,
incomplete. Mesoscutum trapezoidal. Axilla triangular. 1988; van Noort and Rasplus, 1997; Farache et al., 2013).
Scutellum round. Propendeum transverse. The lack of taxonomical knowledge has hampered the
Metasoma. Without petiole. Ovipositor sheath understanding of the ecology and evolution of fig wasps
length 3× longer than body. inhabiting this speciose region.
Male - Head width 0.2–0.6 mm and pear shaped with Given that there are 26 species of native Ficus in
short seta (Fig. 4A). Head longer than wide. Mandible Taiwan, an estimated 70 species of sycoryctine fig
length 0.1–0.3 mm. Eye red, one-seventh of head length. wasps are yet to be discovered. Taxonomical studies of
Body yellow, apterous and body length 2.6–3.5 mm. Taiwanese fig wasps were discontinued in 1999. Only
(Figs. 4B–C). eight species of sycoryctine fig wasp have been
Etymology: The specific name refers to the type of identified in Taiwan. Two new sycoryctine species
locality of Taipei City in Taiwan. associated with F. benguetensis are described and
Host: Reared from syconia of F. benguetensis. illustrated in this paper. The female Philotrypesis can
Biology: This species could be parasitoid fig wasps easily be assigned to Philotrypesini because of the
in the Ceratosolen, Eupristina and Kradibia genera. presence of tubular urotergites of the terminal segments
This species may have some strong seasonal variation in (7th and 8th segments) and its ovipositor. The
abundance and distribution. extraordinarily long ovipositor not only functions as an
159Taiwania Vol. 63, No. 2 Fig. 3. Female Sycorycteridea taipeiensis sp. nov. A: Head frontal view; B: Antenna; C: Lateral view; D: Hind leg; E: Wings; F: Dorsal view of mesosoma. Scale bar = 0.5 mm. 160
June 2018 Wong et al.: New fig wasps associated with Ficus benguetensis
Fig. 4. Male Sycorycteridea taipeiensis sp. nov. A: Head frontal view; B: Dorsal view of mesosoma; C: Lateral view. Scale bar = 0.2
mm.
egg-laying apparatus but is also capable of piercing the phylogeny for the Sycoryctinae reinstated all of the
fig wall. Distinguishing male Philotrypesis is former names. These genera are difficult to
sometimes difficult because of polymorphism; distinguished due to morphological conservatism.
moreover, some resemble male Sycoscapter. Female Sycorycteridea can be differentiated from other
Sycorycteridea was first established by Abdurahiman Sycoryctini species by their anellus number and long
and Joseph (1975), later Bouček (1988) synonymized slender, slightly swollen at the apex of its ovipositor;
Sycoscapter, Sycoryctes, Arachonia, Sycoscapteridea male Sycorycteridea males exhibit a pear-shaped head,
and Sycorycteridea into a single genus Sycoscapter, but which is quite different from the square-shaped male
Segar et al. (2012) in their study on global molecular Philotrypesis.
161Taiwania Vol. 63, No. 2
Although Philotrypesis taida sp. nov. and Gibson, G.A.P. 1997. Morphology and terminology. In:
Sycorycteridea taipeiensis sp. nov. both inhabit the Gibson G. A. P., J. T. Huber and J. B. Woolley (eds.)
same host (Ficus benguetensis), their relationships with Annotated Keys to the Genera of Nearctic Chalcidoidea
each other as well as with pollinating species remain (Hymenoptera): 16-44. NRC Research Press, Ottawa,
Canada.
unclear. To determine the trophic level of these two Jiang, Z-.F., D.-W. Huang, L.-L. Chen, W.-Q. Zhen, Y.-G.
nonpollinating species, a precision-cut tissue slice Fu and Z.-Q. Peng. 2006. Rampant host switching and
technique may be required to acquire evidence of their multiple female body colour transitions in Philotrypesis
feeding habits. Collecting Sycorycteridea taipeiensis sp. (Hymenoptera: Chalcidoidea: Agaonidae). J. Evol. Biol.
nov. is rare because it is sporadically distributed. 19(4): 1157-1166.
Extensive collection of Sycorycteridea taipeiensis sp. Jousselin, E., S. van Noort and J.M. Greeff. 2004. Labile
nov. is therefore encouraged to enhance knowledge male morphology and intraspecific male polymorphism in
about this species. the Philotrypesis fig wasps. Mol. Phylogenet. Evol. 33(3):
706-718.
McLeish, M.J., G. Beukman, S. van Noort and T.C.
ACKNOWLEDGEMENTS Wossler. 2012. Host-plant species conservatism and
ecology of a parasitoid fig wasp genus (Chalcidoidea;
The authors thank Prof. Chiun-Cheng Ko and Mr. Sycoryctinae; Arachonia). PLoS ONE 7(9): e44804.
Chi-Man Leong from the Department of Entomology at Moore, J.C., D.J. Obbard, C. Reuter, S.A. West and J.M.
National Taiwan University for their generous advice and Cook. 2009. Male morphology and dishonest signalling in
suggestions. We also thank Dr. Yuan-Tung Shih for his a fig wasp. Anim. Behav. 78(1): 147-153.
guidance in slide-making techniques for chalcidoid wasps. Murray, M.G. 1987. The closed environment of the fig
This work was supported by the Bureau of Animal and Plant receptacle and its influence on male conflict in the Old
Health Inspection and Quarantine, Council of Agriculture, World wasp, Philotrypesis pilosa. Anim. Behav. 35(2):
Taiwan (106AS-9.5.3-BQ-B1(2)), and Ministry of Science 488-506.
and Technology, Taiwan (104-2311-B-002-031). Noyes, J.S. 1982. Collecting and preserving chalcid wasp
(Hymenoptera: Chalcidoidea). J. Nat. Hist. 16(3): 315-334.
LITERATURE CITED Rasplus, J.Y., C. Kerdelhue, I. Le Clainche and G. Mondor.
1998. Molecular phylogeny of fig wasps Agaonidae are not
Abdurahiman, U.C. and K.J. Joseph. 1975. Three new monophyletic. Evolution 321(6): 517-527.
chalcidoidea (Hymenoptera) from India. Oriental Insects Segar, S.T., C. Lopez-Vaamonde, J.Y. Rasplus and J.M.
9(1): 99-109. Cook. 2012. The global phylogeny of the subfamily
Bain, A., H.-Y. Tzeng, W.-J. Wu and L.-S. Chou. 2015. Sycoryctinae (Pteromalidae): parasites of an obligate
Ficus (Moraceae) and fig wasps (Hymenoptera: mutualism. Mol. Phylogenet. Evol. 65(1): 116-125.
Chalcidoidea) in Taiwan. Bot. Stud. 56(1): 11. Tzeng, H.-Y., L.-J. Tseng, C.-H. Ou, K.-C. Lu, F.-Y. Lu
Bean, D. and J.M. Cook. 2001. Male mating tactics and lethal and L.-S. Chou. 2008. Confirmation of the parasitoid
combat in the nonpollinating fig wasp Sycoscapter feeding habit in Sycoscapteridea, and their impact on
australis. Anim. Behav. 62(3): 535-542. pollinator abundance in Ficus formosana. Symbiosis 45:
Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): a 129-134.
biosystematic revision of genera of fourteen families, with van Noort, S. and J.Y. Rasplus. 1997. Revision of the
a reclassification of species. CAB International, otiteselline fig wasps (Hymenoptera: Chalcidoidea:
Wallingford, UK. 832 pp. Agaonidae), I: the Otitesella digitata species-group of the
Chen, Y.-R., W.-C. Chuang and W.-J. Wu. 1999. Chalcid Afrotropical region, with a key to Afrotropical species of
wasps on Ficus microcarpa L. in Taiwan (Hymenoptera: Otitesella Westwood. Afr. Entomol. 5: 125-147.
Chalcidoidea). J. Taiwan Mus. 52: 39-79. Wang, R., A. Matthews, J. Ratcliffe, L. J. Barwell, Y.-Q.
Chou, L.-Y. and C.-Y. Wong. 1997. New records of three Peng, L.-S. Chou, H. Yu, H.-W. Yang and S. G.
Philotrypesis species from Taiwan (Hymenoptera: Compton. 2014. First record of an apparently rare fig wasp
Agaonidae: Sycoryctinae). Chinese J. Entomol. 17(3): feeding strategy: obligate seed predation. Ecol. Entomol.
182-186. 39(4): 492-500.
Compton, S.G., S. van Noort, M. McLeish, M. Deeble and Yoder, M. J., I. Miko, K.C. Seltmann, M.A. Bertone and
V. Stone. 2015. Sneaky African fig wasps that oviposit A.R. Deans. 2010. Gross anatomy ontology for
through holes drilled by other species. Afr. Nat. Hist. 5: Hymenoptera. PLoS ONE 5(12): e15991.
9-15.
Farache, F.H., A. Cruaud, G. Genson, R.A.S. Pereira and
J.Y. Rasplus 2013. Taxonomic revision and molecular
phylogeny of the fig wasp genus Anidarnes (Hymenoptera:
Sycophaginae). Syst. Entomol. 38(1): 14-34.
162You can also read
